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Fruit and seed biometry and germination of

Victoria amazonica (Poepp.) J.C. Sowerby
(Nymphaeaceae) from the Pantanal...

Article in Acta Scientiarum Biological Sciences October 2016

DOI: 10.4025/actascibiolsci.v38i2.28621

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Doi: 10.4025/actascibiolsci.v38i2.28621

Fruit and seed biometry and germination of Victoria amazonica

(Poepp.) J.C. Sowerby (Nymphaeaceae) from the Pantanal floodplain
Luiz Ricardo dos Santos Tozin1*, Liana Baptista de Lima Corra-da-Costa2 and Edna
Scremin-Dias2
1
Departamento de Botnica, Instituto de Biocincias de Botucatu, Universidade Estadual Paulista Jlio de Mesquita Filho, Cx. Postal 549,
Botucatu, So Paulo, Brazil. 2Laboratrio de Botnica, Centro de Cincias Biolgicas e da Sade, Universidade Federal de Mato Grosso do Sul,
Campo Grande, Mato Grosso do Sul, Brazil. *Author for correspondence. E-mail: ricardo.tozin@gmail.com

ABSTRACT. Numerous plant species are easily established in the wide flood plains of the Pantanal
wetlands due to the environmental heterogeneity. The aquatic macrophytes excel in permanently flooded
areas, particularly Nymphaeaceae species. Victoria amazonica, popularly known as the vitria-rgia, is
hallmarked for its beauty. However, the biology and conditions necessary for the seed germination of this
flowering plant remain unknown. In the present study, the fruit and seed morphology and biometry were
described and the seed germination was evaluated under different abiotic conditions. To this end, mature
fruits of V. amazonica were collected from the bays near Paraguay River in the south Pantanal floodplain.
The fruits and seeds were described and measured using digital caliper. Intact and mechanically scarified
seeds were germinated under different temperature, light and substratum conditions, and the initial
development was described. The fruits measured 67.5 x 119.7 mm in size and contained 100-700 seeds.
The average seed measured 10.6 mm in length and 9.8 mm in width. The highest germination occurred at
25oC, independent of the light condition. The seeds were considered neutral photoblastic. The seedlings
showed heterophylly, and the main root was degenerated, forming adventitious roots. Morphological
differences were observed in seedlings developed under different light conditions.
Keywords: aquatic macrophyte, hydrophytes, water lilies, vitria-rgia.

Biometria do fruto e da semente e germinao de Victoria amazonica (Poepp.) J.C. Sowerby

(Nymphaeaceae) do Pantanal
RESUMO. O Pantanal, ampla plancie inundvel, facilita o estabelecimento de inmeras espcies vegetais
por sua heterogeneidade ambiental. As macrfitas aquticas se destacam em reas permanentemente
alagadas, com destaque para a famlia Nymphaeaceae. A espcie Victoria amazonica, conhecida popularmente
como vitria-rgia, marcada por sua beleza. Entretanto, sua biologia e as condies necessrias
germinao de suas sementes permanecem desconhecidas. Nesse estudo foram descritas a biometria e a
morfologia do fruto e da semente, e avaliada a germinao em diferentes condies abiticas. Para isso,
frutos de V. amazonica foram coletados em baias prximas ao rio Paraguai no Pantanal sul. Os frutos e as
sementes foram descritos e mensurados usando paqumetro digital. Sementes ntegras e escarificadas
mecanicamente foram submetidas a testes de germinao em diferentes condies de temperatura, luz e
substrato, e o desenvolvimento inicial das plntulas foi descrito. Os frutos mediram 67,5 x 119,7 mm, e
portavam 100-700 sementes. As sementes mediram 10,6 mm de comprimento e 9,8 mm de largura em
mdia. A maior porcentagem de germinao ocorreu a 25oC independente da condio de luz. As sementes
foram consideradas fotoblsticas neutras, e as plntulas possuem heterofilia, degenerao da raiz principal e
formao de razes adventcias. Foram observadas diferenas morfolgicas nas plntulas desenvolvidas na
presena e na ausncia de luz.
Palavras-chave: macrfita aqutica, hidrfitas, vitria-rgia.

Introduction lead to the significant accumulation of sediment and

The Pantanal floodplain shows diverse aquatic
vegetation, which occupies periodically or
permanently flooded areas (Allem & Valls, 1987).
The lakes receive water from rivers through
drainage areas (Bacani & Sakamoto, 2007), similar to
all wet areas on the planet. These water dynamics
seeds from several plant species (Van Der Valk &
Davis, 1976).
The macrophyte populations, as determined by
seed banks, are efficient in recovering communities
after flooding or drought (Ferreira, Mormul,
Thomaz, Pott, & Pott, 2011). Seasonal flooding

Acta Scientiarum. Biological Science Maring, v. 38, n. 2, p. 221-227, Apr.-June, 2016

222
favors the occurrence of species adapted to flooding
during the wet season and drought during the dry
season (Scremin-Dias, Lorenz-Lemke, & Oliveira,
2011). In particular, macrophytes stand out in
permanently flooded areas because these plants
show peculiar adaptations that facilitate growth at
different moisture gradients (Esteves, 1998).
The Nymphaeaceae is quite representative
among aquatic macrophytes occurring in Pantanal
areas, with seven Nymphaea species and one Victoria
species (Pott & Pott, 2000). Victoria amazonica
(Poepp.) JC Sowerby is a floating fixed hydrophyte
with peltate and circular leaves of up to 2 meters in
diameter that are connected to the main stem
through long flexible secondary stems (Sculthorpe,
1967; Cronk & Fennessy, 2001). Previous studies
have focused on the floral morphology (Corra,
1952), blooming and pollination processes
(Seymour & Matthews, 2006), stamen morphology
(Heinsbroek & Heel, 1969), and ovary
pseudosyncarpia (Weberling, 1992) of these plant
species. In addition, specific information about the
flower, fruits, seeds and seedlings morphology of
plants collected from the Brazilian Amazon has been
reported (Rosa-Osman, Rodrigues, Mendona,
Souza, & Piedade, 2011). However, little is known
about the conditions for the seed germination of
V. amazonica and the morphological characteristics of
the initial development of these plants. Does the
occurrence of this species in the Pantanal floodplain
(Pott & Pott, 2000), where the climatic
characteristics differ from the Amazon, might reflect
distinct seed and seedling peculiarities and the
adaptation of these plants to this environment?
Successful seedling establishment in this
environment depends on the seed germination
process. Germination is the stage in the plant life
cycle that directly influences plant distribution in the
environment (Souza, Pacheco, Matos, & Ferreira,
2007). The maximum germination percentage was
observed in germination tests performed under
optimum conditions for each species (Ministrio da
Agricultura, Pecuria e Abastecimento [MAPA], 2009),
and several factors affect germination. Thus,
identifying the optimum temperature and light
conditions for maximum germination is essential for
propagation of certain species. In addition, this
knowledge provides information concerning species
biology and adaptation to the environment and
provides subsidies for preservation of these plants.
In the present study, we described the fruit and
seed morphology and biometry, and evaluated the
seed germination of Victoria amazonica under
different abiotic conditions. This study provides the
Acta Scientiarum. Biological Science
Tozin et al.
first information about the fruit, seeds and seedlings
of V. amazonica population occurring in the Pantanal
floodplain.
Material and methods
Study area and seeds processing
The Pantanal floodplain in Mato Grosso do Sul
show a tropical climate Aw, according to Kppen,
with a 1.070 mm average annual rainfall. The rainy
season occurs from October to March, and the dry
season occurs from April to September (Soriano,
1997). The average annual temperature is 26C. In
addition, frost might sporadically occur (Cadavid
Garca, 1984).
The fruit samples were collected in October
2010 from three bays under Paraguay River
influence (17 58 19.8 S and 57 29 28.8 W; 17
54 41.8 S and 57 27 37.5 W; 18 0.2 40.0 S and
57 29 33.0 W) in Corumb, Mato Grosso do Sul
State, Brazil. In the total, 23 mature fruits were
collected, and vouchers were deposited in the
Herbarium CGMS, Universidade Federal de Mato
Grosso do Sul (UFMS), Campo Grande, Brazil,
under registration 29788.
The fruits were transported in Styrofoam boxes
to the Seed Ecophysiology Laboratory. To
determine the initial water content, the seeds were
immediately extracted from the fruits, and the aryl
was removed.
Fruit and seed morphology and biometry
The fruit and seed morphology were analyzed in
field and in laboratory with stereomicroscope Leica,
and described according to Hoehne (1948), Judd,
Campbell, Kellogg, Stevens and Donoghue (2002)
and Vidal and Vidal (2003) terminology.
The biometrics data and number of seeds per
fruit were determined for the 23 fruits collected.
Biometrics data were determined for 400 seed
samples. The fruits and seeds were measured using a
digital caliper. The fruit length was considered from
the peduncle insertion to the base of the perianth,
and fruit width was obtained perpendicular to this
area. The seed length was considered from the hilo
to the opposite side, and seed width was obtained
perpendicular to this area. The seed moisture
content was determined through oven drying at
105C for 24 hours, according to MAPA (2009). The
1000 seeds weight was obtained from.
Germination and seedling morphology
The seeds were distributed into paper towel rolls
moistened with 3.0 times its weight in water and
maintained in a growth chamber BOD equipped
Maring, v. 38, n. 2, p. 221-227, Apr.-June, 2016
Morphology and germination of vitria-rgia 223

with four fluorescents light of 250 v until stable 1B-C); without the white bag the seed shows dark
germination was obtained. The temperatures brown color (Figure 1D). They are placed along the
conditions included 20, 25, 30 and 20-30C main axis, featuring axial placentation, and to
(12 hours at 20C and 12 hours at 30C). At all develop involved in pulp. The embryo has two
temperatures, the experiment was conducted in the cotyledons, one developed and the second stunted.
absence and presence of light to evaluate the effect The embryo is placed in the hilar region,
of this factor on germination. The seeds were surrounded by whitish reserve tissue. The length of
considered germinated upon primary root emission the seeds was 10.6 ( 4.0) mm, and the width was
of at least 3 mm. The seeds were submitted to 9.8 ( 0.6) mm. The thousand seed weight was
mechanical scarification of the seed integument 392.9 ( 11.5) g. The water content in the seeds
using sandpaper number 3 in the hilum opposite immediately after extraction from the fruits was
area. These seeds were placed on paper towel 44.8%.
moistened with 3.0 times its weight in water, kept at
25, 30 and 20-30C (12 hours at 20C and 12 hours
at 30C). All treatments were performed using four
repetitions with 40 seeds each.
To establish conditions similar to the natural
environment of the species occurrence, 40 seeds
were distributed into four trays filled with washed
sand and 40 seeds in four trays containing soil
collected at the area of the species occurrence. The
seeds were placed at 2 cm deep. The set was
moistened at a water depth of approximately 1 cm
and maintained at room temperature (25 2C)
under laboratory conditions and constant light.
The results were expressed as a germination
percentage, and efficient germination was
considered to be a higher germination percentage
obtained in a shorter period of time.
The experiment was designed in a factorial 4 x 2
(temperature and light condition) and 3 x 2
(temperature and scarification). We used a
completely randomized experimental design, and
the data were analyzed using ANOVA, followed by
Tukeys test at 5% probability.
The seedling morphology was described
according to Souza (2009) and Wheeler Haines
(1975).
Figure 1. Fruit, seed and seedling morphology of Victoria
Results amazonica (Poepp.) JC Sowerby. A. General aspects of fruit and
perianth in decomposition (asterisk). Arrow indicates the seed
Fruit and seed morphology and the white pulp. B. General aspects of seeds. C. Detail of the
The fruits are covered by spine, and have seed surrounded by a white bag, probably aryl. D. Mature seed
without aryl. E. Seedlings developed in the absence of light.
greenish color when immature and brown color at Arrow indicates the developed mesocotyl. F. Seedlings developed
maturity. They are polismermic, fleshy, indehiscent, in the presence of light. Scale bars: A= 2 cm; B, E-F = 1 cm;
apocarpic, multicarpellate, plurilocular and berry C = 5 mm; D = 3 mm.
type. In the apical portion is placed the perianth in
Germination
decomposition (Figure 1A). They develop
underwater and at maturity the fruit rots releasing Germination was slow and non-uniform, once
the seeds. The fruit average size was 67.0 ( 13.1) the first seeds germinated at 60 days after treatment
mm in length and 119.7 ( 19.5) mm in width. The was initiated, and germination was observed for
number of seeds per fruit ranged 100 - 700, with an approximately 170 days. At 25C promoted higher
average of 300 to 400 seeds per fruit. percentage germination, regardless of the lighting
The seeds are hard, globular shape and conditions. At 30C, germination ranged from 16.25
surrounded by a white bag, probably aryl (Figure to 18.75%, with statistically similar results to those

Acta Scientiarum. Biological Science Maring, v. 38, n. 2, p. 221-227, Apr.-June, 2016

224 Tozin et al.

obtained for the best germination temperature. successful for seedling emergence from both non-
Temperatures of 20 and 20-30C were inadequate, scarified and scarified seeds.
showing no success in germination (Table 1).
Seedling morphology
Table 1. Germination percentage (%) of Victoria amazonica
(Poepp.) JC Sowerby seeds under different lighting conditions The seedling development was irregular, occurring
(light and dark) and temperatures. for 170 days. The embryos showed two cotyledons,
Germination percentage developed and stunted, in the hilar region surrounded
Temperature
Light Dark by a whitish reserve tissue. The first step in the seed
20C 0.00 0.00 Ba 0.00 0.00 Ca germination process is the embryonic axis protrusion,
25C 25.60 5.90 Aa 25.62 2.37 Aa
30C 18.75 5.95 Aa 16.25 5.95 Ba starting the seedling stage. At this stage, the protrusion
20-30C 1.00 0.00 Ba 1.00 0.00 Ca of the acicular cotyledon occurs simultaneously with
C.V. 17.1
Capital letters indicate comparisons between lines, and lowercase letters indicate
hypocotyl emergence. The primary root emerges from
comparisons between columns. Means followed by the same letter do not differ the basal region of the hypocotyl and is subsequently
according to Tukey test at 5% significance. C.V.: coefficient of variation.
replaced with adventitious roots arising from the
The germination of the scarified seeds was slow cotyledon node. The acicular cotyledon gradually
and non-uniform, once the first seeds germinated at degenerates after the onset of the first sagittate eophyll.
70 days after experiment was initiated, and stabilized Morphological differences between seedlings
at 150 days after. Compared with non-scarified developed in the absence (Figure 1E) and presence of
seeds, scarified seeds showed optimized germination light (Figure 1F) were observed. In seedlings emerging
at 20-30C in the dark, but low germination was in the absence of light, the mesocotyl extends below
observed at the other temperatures at 25 and 30C in the cotyledonary nodes of up to 30 mm in length,
both light condition (Table 2). connecting seedling to seed reserves (Figure 1E). In
After 170 days, seedling emergence from non- seedlings obtained in the presence of light, this
scarified and scarified seeds was 24.00% ( 5.00) structure is a maximum of 3 mm in length (Figure 1F).
and 22.64% ( 4.32), respectively, in soil obtained The Figure 2 shows the evolution of germination
from the collection site. Washed sand was not process in the presence of light.
Table 2. Germination percentage (%) of scarified and non-scarified Victoria amazonica (Poepp.) JC Sowerby seeds under different lighting
conditions (light and dark) and temperatures.
Light Dark
Temperature
Non-scarified Scarified Non-scarified Scarified
25C 25.60 5.90 Aa 16 4.50 Ab 25.65 2.37 Aa 12 2.55 Ab
30C 18.75 5.95 Aa 2 0.50 Bb 16.25 5.95 Ba 13 2.30 Aa
20-30C 1.00 0.00 Bb 8 1.50 ABa 1.00 0.00 Cb 13 3.45 Aa
C.V. 24.9 14.6
Capital letters indicate comparisons between lines, and lowercase letters indicate comparisons between columns, Means followed by the same letter do not differ according to Tukey
test at 5% significance. C.V.: coefficient of variation.

Figure 2. Evolution of germination process of Victoria amazonica (Poepp.) JC Sowerby in the presence of light. Scale bar = 2 cm.

Acta Scientiarum. Biological Science Maring, v. 38, n. 2, p. 221-227, Apr.-June, 2016

Morphology and germination of vitria-rgia 225

Discussion germinated only when scarified, indicating the

importance of this in nature, when seeds are
The germination percentage of V. amazonica
subjected to temperature variations between day and
seeds was low, and this process was irregular and
night. Frequently, hard impermeable seed tegument
slow. The first seeds germinated after two months,
is typical of physical dormancy, and scarification
and germination continued for approximately six
methods may be use for overcome this dormancy
months, suggesting the presence of dormancy. In
and accelerate germination (Marcos Filho, 2005).
addition, germination occurred independent of the
Scarification can occur in nature when the seed
light conditions, but the temperature and substrate
passes through the digestive tract of an animal or is
affected germination and were limiting under some
naturally roughened by rain and substrate friction.
conditions.
Although V. amazonica seeds present a hard
The fruit develops underwater environment due
tegument, scarification did not promote higher
to flower stalk curved in spiral-shaped after
germination rates, suggesting that the seed has
pollinated (Sculthorpe, 1967). The fruit permanence
another type of combined dormancy that is likely
underwater, allows the pericarp decomposition, and
associated with the immature embryo and/or the
the organic material can be used as food by
presence of inhibitor hormones. Notably, failed
underwater fauna. This process makes seeds
germination of V. amazonica seeds in washed sand
exposure on the water environment; and the
observed in the present study requires further
presence of aryl air bag shaped allow the seeds float
examination.
at the water surface and be dispersed. The funicle
In the present study, V. amazonica seeds were
aryl has already been described as efficient on the
considered neutral photoblastic and germinated
seeds dispersion, since its involves all the seed,
independent of the light conditions. However, seeds
allowing its fluctuation (Barroso, Amorim, Peixoto,
of the same species from Amazon populations were
& Ichaso, 1999). The seeds can penetrate the
negative photoblastic, germinating only in the total
adjacent sediment and germinate under ideal light,
absence of light (Rosa-Osman, 2010). Ferreira and
oxygen and space for growth situation (Cronk &
Borghetti (2004) considered V. amazonica to be
Fennessy, 2001). The fruit and seed morphology of
recalcitrant and dormant seeds because they have
V. amazonica occurring in the Pantanal floodplain do
inhibitory substances in the aryl besides hard
not differ from individual occurring in the Amazon
tegument. Crawford (1992) reported that Victoria
(Rosa-Osman et al., 2011).
seeds lose viability when subjected to drying.
Low germination percentage described here does
Preliminary tests performed in the present study,
not represent a standard for this plant family.
using clean seed samples, scarified and soaked in
Indeed, Nymphaea odorata showed 100% germination
tetrazolium solution at 1%, the embryo showed red
(Richards & Cao, 2012). However, for V. amazonica
staining after 8 hours of soaking (data not shown),
seeds from the Amazon, the germination percentage
indicating that 100% of seeds were viable.
was also low and only occurred under hypoxic
The degeneration of the main root during the
conditions, total light absence and mechanical
first week of seedling formation and the emergence
scarification (Rosa-Osman, 2010). Mitra, Basu, &
Mukhopadhyay (1982) placed V. amazonica seeds on of adventitious roots in the cotyledon node are
moistened cotton in a greenhouse with temperatures characteristics described for monocots embryos and
ranging from 22.8 to 35.9C, and observed 22% seedlings (Pereira, Pereira, Rodrigues, & Andrade,
germination after 4 months. The data obtained in 2008; Nakamura & Scatena, 2009). Although this
the present study suggest that among the evaluated species belongs to the basal angiosperms, this family
conditions, 25C is the best condition for obtaining is closely related to the monocot group, where this
V. amazonica seedlings in the Pantanal floodplain, feature is common (Angiosperm Phylogeny Group
regardless of the light conditions, thereby extending [APG] III, 2009). Mitra et al. (1982) also described
the results of a previous study (Mitra et al., 1982). the occurrence of taproot degeneration for V.
Mechanical scarification was not efficient to amazonica seedlings obtained from the Botanical
increase the percentage of V. amazonica seed Garden lakes in India.
germination from the Pantanal floodplain, as the Four distinct eophylls have been described for
highest percentage germination was observed for the V. amazonica seedlings from the Amazon (Rosa-
non-scarified seeds. However, V. amazonica seeds Osman et al., 2011), but this work was possible to
from the Amazon showed successful germination describe until the formation of second eophyll.
only under conditions scarification (Rosa-Osman, Slightly more than one week after the protrusion of
2010). Moreover, seeds maintained at 20-30C the embryonic axis, we observed seedlings with the

Acta Scientiarum. Biological Science Maring, v. 38, n. 2, p. 221-227, Apr.-June, 2016

226
first eophyll formed and the second eophyll in
formation. The heterophylly occurs in
Nymphaeaceae species only in the seedling stage
(Hoehne, 1948; Sculthorpe, 1967); however, in the
laboratory, it was not possible to verify the
emergence of all eophylls because seedlings did not
develop until the next adulthood. The continued
development would only be possible with seedling
transplanted under ideal conditions of vegetative
growth and development, in soil with nutrients and
enough light for photosynthesis. The development
of the seeds on paper towels was not continued
because the reserve tissue was not sufficient to
support the development and training of other
leaves. Although germination was performed in the
laboratory, we attempted to cultivate the seedlings
obtained in the germination test, but this endeavor
was not successful, reflecting a lack of basic
information on the needs of this species for
vegetative growth and the small number of seedlings
and appropriate structures to maintain these plants
under different experimental conditions.
The presence of the mesocotyl in seedlings
obtained in the dark has been previously described
as a type of underground stem (Rosa-Osman, 2010).
However, Wheeler Haines (1975) identified this
structure in other Nymphaeaceae species as a
mesocotyl. The mesocotyl is the first internode
connection between the coleoptile and the plumule.
This characteristic is common in Poaceae and
Cyperaceae species, and although it has evolved
independently in each group, this feature would be
important to characterize these families as monocots
(Wheeler Haines, 1975).
The presence of a functional mesocotyl, in the
absence of light, might reflect an important adaptive
character for V. amazonica because this structure
facilitates successful seedling establishment and
germination below the soil and water surface.
Mesocotyl elongation facilitates the elevation of the
apical meristem to the soil surface, and with the
emergence of photosynthetically active eophylls
above the surface, assimilation is initiated with a
completely independent seed reserve thereafter.
Despite the few studies available on V. amazonica
seeds, these data are consistent with the indication that
the low percentage of germination obtained for this
species is a feature reflecting high seed production,
given the predominance of the number of seeds,
ranging from 300 to 400 seeds, with large sizes.
However, because the non-germinated seeds showed
no fungal contamination or morphological changes, it
is likely that germination continues after the 170 days
evaluated in this study. This slow germination could
represent an important contribution to the successful
Acta Scientiarum. Biological Science
Tozin et al.
establishment of this species, using dormancy as a
germination guarantee over time, increase the
possibility of seedling recruitment under different
environmental conditions, and increase its
environmental occurrence in areas where water
seasonality is intense and unpredictable (Scremin-Dias,
2000; Pott, Oliveira, Damasceno-Junior, & Silva, 2011;
Scremin-Dias et al., 2011). Moreover, long mesocotyl
formation in seedlings germinated in the dark might
represent an important role under natural conditions
because the substrate on which the seeds germinate
under natural conditions is dark and muddy,
preventing the entry of light.
Conclusion
The germination percentage of V. amazonica seeds
was low, and this process was irregular and slow. The
highest germination occurred at 25oC, independent of
the light condition. The first seeds germinated after
two months, and germination continued for
approximately six months, suggesting the presence of
dormancy. The seedlings showed heterophylly, and the
main root was degenerated, forming adventitious roots.
Morphological differences were observed in seedlings
developed under different light conditions.
This is the first experimental approach on the seed
germination of V. amazonica occurring in the Pantanal
floodplain. From this study new questions were asked,
as how substrate and oxygenation affect in the seed
germination, and the factors leading to break
dormancy. Further studies are being conducted to
increase knowledge about these aspects.
Acknowledgements
The authors would like to thank the Conselho
Nacional de Desenvolvimentos Cientfico e Tecnolgico
[CNPq] by the scholarship granted to L.R.S. Tozin
(Pibic 2010/2011), the staff of the Laboratory of
Botany, UFMS, Campo Grande, for assistance with
the sample preparation; and the Thales Dias Leandro
by the support in the field areas.
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Received on July 22, 2015.
Accepted on April 5, 2016.
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Maring, v. 38, n. 2, p. 221-227, Apr.-June, 2016